Opinion TRENDS in Parasitology Vol.22 No.

11

Competitive displacement in
Triatominae: the Triatoma infestans
success
Marcos H. Pereira1, Nelder F. Gontijo1, Alessandra A. Guarneri2, Maurı́cio R.V.
Sant’Anna3 and Liléia Diotaiuti4
1
Departamento de Parasitologia do Instituto de Ciências Biológicas da Universidade Federal de Minas Gerais,
Caixa Postal 486, 31270-901, Belo Horizonte, MG, Brazil
2
Departamento de Microbiologia e Parasitologia da Universidade Federal de Santa Catarina, Caixa Postal 476, 88.040-900,
Florianópolis, SC, Brazil
3
Molecular & Biochemical Parasitology Group, Liverpool School of Tropical Medicine, Pembroke Place, Liverpool, L3 5QA, UK
4
Centro de Pesquisas René Rachou/Fiocruz, Caixa Postal 1743, 30190-901, Belo Horizonte, MG, Brazil

Brazil has just been certificated by Pan American Health
Organization as ‘free of Chagas disease transmission
due to Triatoma infestans’. During the early 1980s, this
species of blood-sucking bug alone was considered
responsible for approximately 80% of Chagas disease
transmission. But it was not always so. The species
originally abundant in houses of central and eastern
Brazil was Panstrongylus megistus, which seems to
have been progressively displaced from houses by
T. infestans during the past century. Indeed, T. infestans
seems able to displace other Triatominae in artificial
environments. Recent studies suggest that it might
simply be because T. infestans feeds more efficiently
than its Triatominae competitors.
(J.C.P. Dias, PhD thesis, Federal University of Minas
Gerais, 1982) [4]. Many of the earlier studies reported
the co-existence of T. infestans and P. megistus in Minas
Gerais [5–8], although in most of these, T. infestans pre-
sented greater densities. As T. infestans continued its
northern dispersal in Brazil, reaching its greatest
dispersion in 1981 [9] (Figure 1), it seemed to displace
not only P. megistus but several other Triatominae such as
T. sordida in central Brazil [10], and T. brasiliensis and
T. pseudomaculata in parts of the northeast of Brazil. [9].
In those states where T. infestans was most abundant,
Chagas disease prevalence exceeded the national average
of 4.2%: in Minas Gerais (8.8%), Rio Grande do Sul (8.8%),
Goiás (7.4%) and Bahia (5.4%) [11].

Historic perspective
Why was T. infestans so successful in displacing
Current theory holds that the blood-sucking bug Triatoma
other Triatominae in Brazil?
infestans originated from the Andean valleys of Bolivia,
Population dynamics and intraspecific competition
where sylvatic populations can still be found among rock
T. infestans, as well as other triatomine bugs (Hemiptera,
piles associated with wild rodents [1]. Its original domes-
Triatominae), are hematophagous insects in both adult
tication in that region might have been associated with the
and immature (first to fifth instars nymphs) stages and
hunting and domestication of wild guinea-pigs by pre-
their principal hosts are birds and mammals.
Columbian Andean cultures, but its subsequent dispersal
Because Triatominae are blood-feeding hemimetabolous
through Argentina and Paraguay into southern Brazil is
insects, their life cycles and population dynamics depend
believed to have been in association with human migra-
primarily on their interactions with vertebrate hosts. In a
tions during the later part of the 19th century [2]. Ento-
domestic habitat, the number of triatomine bugs is related
mological surveys following the work of Carlos Chagas in
to the number of occupants because the nutritional status of
the early 1900s give no evidence for the presence of T.
the bugs depends on the number of insects per host [12,13].
infestans in the central Minas Gerais state of Brazil. At
The mean amount of blood taken by T. infestans and
that time, in Minas Gerais, T. infestans was present only in
P. megistus from non-anesthetized hosts is inversely related
the south, near the border with São Paulo state [3].
to insect density [13,14]. Increased insect density causes an
The scale of dispersal of T. infestans became evident
increased perception of bites by the host, which probably
following studies carried out in Bambuı́ (a mid-western
diminishes blood-meal size because of more frequent
region of Minas Gerais), where Panstrongylus megistus
interruptions in feeding [15]. Such a reduction in blood
had previously been described as the main vector of Chagas
intake leads to a prolonged nymphal development time,
disease. Although the first record of T. infestans in Bambuı́
reduced egg laying and increased dispersal by adults. These
was only in 1939, by the early 1940s T. infestans repre-
factors combine to regulate insect population density [16]. A
sented >90% of the triatomines collected inside houses
comparative study using non-anesthetized mice as hosts
Corresponding author: Pereira, M.H. (marcoshp@icb.ufmg.br). showed that T. infestans can ingest larger blood meals than
Available online 12 September 2006. P. megistus at different densities (Figure 2). This better
www.sciencedirect.com 1471-4922/$ – see front matter ß 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.pt.2006.08.012
 Opinion TRENDS in Parasitology
Figure 1. A map of the geographical distribution of Triatoma infestans taken from
entomological surveys between 1975 and 1983 by the Chagas Disease Control
Program (PCDCh). São Paulo state was not included in the entomological surveys.
Abbreviations of the Brazilian states: BA, Bahia; MG, Minas Gerais; RS, Rio Grande
do Sul; SP, São Paulo. Source: PCDCh/Sucan.
exploitation of blood resources might explain why
T. infestans reaches greater densities than P. megistus
inside human dwellings [17].
Interspecific competition
The life cycle of T. infestans is shorter than that of P.
megistus, and it needs smaller amounts of blood [18]. Such
characteristics could provide T. infestans with a significant
competitive advantage over P. megistus if the two species
established populations with similar numbers of
Figure 2. The exploitation of the blood resource (weight gain) at different
population densities after 24 h contact of third instar nymphs of Triatoma
infestans (circles) and Panstrongylus megistus (squares) with non-anesthetized
mice. The data were obtained from linear regressions of mean weight gain (mg)
versus density from Ref. [17]. Bold lines correspond to experimental data and
dotted lines correspond to estimates.
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Vol.22 No.11 517
individuals in a single house. Thus, T. infestans could
displace P. megistus in areas where P. megistus already
existed in stabilized domestic colonies. The colonization by
T. infestans of houses already infested with indigenous
triatomines was probably initiated by a few individuals
transported passively in human belongings [19]. No direct
interference has been observed between the two species
when they have been reared together in the laboratory in
the same recipient [20]. Peak activity of both species occurs
within a few hours of nightfall [21] (H.H.R. Pires, unpub-
lished) so competition between them might involve
interactions with the host, as described above for intraspe-
cific competition.
Feeding behaviour
Triatomine bugs are vessel feeders, obtaining their blood
meals directly from the blood vessels (venules or arterioles)
of their vertebrate hosts. After the bug pierces the host
skin, a probing period characterized by rapid whip-like
intradermal movements of the maxillae can be observed.
Once a suitable vessel is found, probing ceases and the
bugs engorge [22]. The ingestion of blood through the food
canal is aided by the cibarial pump, the movements of
which are regulated by a complex of strong muscles occu-
pying much of the head of the insect [23]. Parameters such
as the negative pressures produced by the cibarial pump,
the dimensions of the food canal, blood viscosity, and the
size of host red cells and their capacity to deform can
influence the ingestion rate [24].
Smith and Friend [25] created the standard technique
for studying feeding behaviour in triatomines, on the basis
of changes in electrical resistance between the insect and
its feeding sources. Smith [26] and Guarneri et al. [27]
improved this technique to record signals generated by the
cibarial pump musculature, similar to an electromyogram.
Further modifications of the technique enabled the feeding
behaviour of early developmental stages to be recorded
because contact involved placing the insect on a metal
mesh rather than implanting or fixing electrodes to the
thorax [28].
Total ingestion rate
On the basis of the electrical signals methodology, several
studies have demonstrated total ingestion rate (TI) to be
the main parameter that determines contact time between
a particular triatomine species and an immobilized host. TI
values for fifth instar nymphs of the species studied to date
vary from 3.2 mg/min for Rhodnius neglectus feeding on
mice to 25.3 mg/min for T. infestans feeding on pigeons
[27,29]. The highest TI values for feeding on mice are
observed for T. infestans (13.8 mg/min) and R. prolixus
(11.9 mg/min), which are the most important vectors of
Chagas disease in South and Central America, respectively
[27,29]. TI also varies considerably between different nym-
phal instars of the same species, for example, from 0.4 mg/
min for first instars to 10.6 mg/min for fifth instars of
T. brasiliensis feeding on humans [28].
Contact time versus total ingestion rate
Contact time with the host for nymphs of T. infestans,
T. brasiliensis and T. pseudomaculata is strongly
518 Opinion TRENDS in Parasitology Vol.22 No.11
correlated (r2 = 0.92) with the ratio between the initial
weight after moulting and the total ingestion rate (IW/
TI), which represents the time an insect requires to double
its body weight by feeding on a specific type of host
(Figure 3). For these species, the initial weight provides
a good estimate of the size of the blood meal for the nymphs
and is also a good estimate of the insect biomass, regard-
less of species or nymphal stage. The contact time is
strongly correlated with the IW/TI ratio (Figure 3) so
triatomines with higher IW/TI values will have more
difficulty in maintaining a good nutritional status as popu-
lation density increases because of their need for a longer
contact time to complete the blood meal. The IW/TI values
for these three species were lower for feeding on pigeons
than on mice. First- and second-instar nymphs of T. bra-
siliensis fed more efficiently than other instars on human
hosts. The same theory could be applied to explain differ-
ences between males and females [28]. A lower IW/TI ratio
for females (10.8) would enable them to feed more effi-
ciently than males (13.9), which might be an adaptation to
increase their fitness in situations of nutritional stress.
This theory is in agreement with observations that males
have systematically lower qualitative nutritional status
than females in T. infestans peridomestic populations from
rural northwestern Argentina [30]. The reduced size of
insects in high-density situations [10,31] might reflect a
selection for individuals with a low IW/TI ratio that can
feed more rapidly.
Factors affecting total ingestion rate
Monitoring the electrical activity of the cibarial pump to
study triatomine feeding behaviour has enabled the total
ingestion rate (TI) to be correlated with other parameters
such as cibarial pump frequency (F), cibarial pump
volume (QLC), contact time (CT) and non-ingestive time
Figure 3. The correlation (r2 = 0.92) between the contact time of Triatominae instar nymphs with their vertebrates hosts and the IW:TI ratio (initial weight: total ingestion
rate). Triatoma infestans, circles; Triatoma brasiliensis, squares; Triatoma pseudomaculata, triangles. Each point is labeled with the nymphal instar (1st to 5th) and the
vertebrate host used as the blood meal source (human, mouse or pigeon). Adapted, with permission, from Ref. [28].
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Box 1. Parameters of feeding behavior
Contact time [CT (min)] is defined as the time during which the
mouthparts of the insect are inserted into the host skin. Total
ingestion rate [TI (mg min1)] is calculated by dividing the weight
gain (mg) by the total contact time (min). Cumulative probing time
(PT) is the period from the insertion of the mouthparts into the host
to the initiation of the engorgement phase. If the bug ends an initial
probing and restarts another elsewhere, the first time is added to the
second and so on successively; non-ingestive time [NIT (min)] is
defined as the period when insects are not pumping, thus
comprising probing time plus any interruption to feeding. The
quantity of liquid ingested per cibarial contraction [QLC (ml
contraction1)] is obtained by dividing the weight gain by the total
number of cibarial pump contractions during the feeding process,
considering blood and artificial diet densities (r) as 1 mg ml1. Pump
frequency [F (contractions min1)] represents the number of cibarial
pump contractions divided by the ingestive time (the time in which
the pump is effectively working).
These variables are linked by Equation I:
TI ¼ QLC  F  r  ½ðCT  NIT Þ=CT  [Eqn I]
(NIT: probing time plus interruptions during engorge-
ment; see Box 1).
The analysis of these parameters during feeding by
different triatomine species on pigeons, mice, humans or
artificial feeders has provided enough information to know
whether differences were because of the vertebrate host,
the triatomine species or the developmental stage [27–29].
The QLC and maximum pumping frequency (obtained by
using artificial feeders) are related to the intrinsic char-
acteristics of the mouthparts of the insect (morphology). In
the triatomine species studied, QLC ranges from
25 nl in
T. pseudomaculata to 100 nl in T. infestans. The QLC
increases more gradually than the body weight and this
asymmetrical growth is responsible for the increase in the
 Opinion TRENDS in Parasitology
IW/TI ratio observed during nymphal development
(Figure 3). There is a remarkable sexual dimorphism of
the QLC, with that of the females being greater than that of
males.
Parameters such as F and NIT are influenced by the
host physiology. Most of the triatomine species feeding on
mice have a lower F value than those feeding on pigeons.
The NIT values are variable. However, except for R. nasu-
tus, the triatomines feeding on mice usually have a longer
NIT. These facts might be related to hemostatic features of
birds and mammals. Birds have thrombocytes that per-
form a similar function to mammalian platelets but are
less effective. They also seem to lack some coagulation
factors, particularly in the intrinsic system [32].
Experiments using salivarectomized R. prolixus showed
that insects had increased difficulty in obtaining a blood
meal from rabbits, as well as a prolonged probing time [33].
Although the probing time is reduced (usually < 10% of the
contact time), it should be important in triggering the host
responses. The insertion of the mouthparts, the move-
ments of the maxillae and saliva deposition within the
skin of the host lead to tissue damage, which is responsible
for liberating mediators and exposing molecules that trig-
ger hemostasis, inflammation and immunological reac-
tions [34,35]. Saliva is continuously liberated during the
feeding process [36] and has an important role in finding
blood vessels during probing and keeping blood flowing
through the feeding canal [37,34]. Comparative studies
showed qualitative and quantitative differences in Triato-
minae salivary antihemostatic activity [38,39]. Activities of
the apyrase enzyme and vasodilator, but not anticlotting
agents, correlated with feeding efficiency of triatomines on
rat [39]. Besides facilitating blood feeding, triatomine
saliva inhibits the generation of nervous impulses, func-
tioning in a similar way to local anaesthetics. This provides
further confirmation of the importance of host perception
in triatomine feeding success [40].
In summary, the mechanical and salivary characteris-
tics of the insect and the host physiology determine the
feeding performance of triatomines. The fact that T. infes-
tans feeds more easily on birds could explain why bugs
collected in goat and pig corrals had a lower qualitative
nutritional status than those captured in chicken coops in
Argentina [30].
The importance of feeding performance (IW/TI)
On human hosts, the fifth instar nymph of T. infestans has a
lower IW/TI ratio (4.1) than P. megistus (7.6; S.E. Barbosa
et al., unpublished) and T. brasiliensis (6.3). A lower IW/TI
ratio means a proportionally greater amount of blood
ingested in the same contact time with the host. This would
explain epidemiological findings in which T. infestans not
only succeeds in establishing larger colonies but also dis-
places P. megistus and T. brasiliensis from human dwellings
when it is introduced in areas where the other two species
were already established. The existence of competition
events among these species is supported by the findings
that P. megistus [41] and T. brasiliensis (A.N. Ramos Jr, MSc
thesis, Federal University of Rio de Janeiro, 2001) recolo-
nized houses from sylvatic specimens after the elimination
of T. infestans (a species only found in domestic and
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Vol.22 No.11 519
peridomestic environments in Brazil). Studies under
laboratory conditions also showed competitive exclusion
of T. sordida by T. infestans [42] that probably could be
explained by a lower IW/TI ratio characteristic of T.
infestans.
Another implication of the better exploitation of blood
resources by Triatominae is related to the dynamics of
defecation and, consequently, the transmission of Trypa-
nosoma cruzi. The defecation timing depends on both the
triatomine species and the bloodmeal size because bugs
with large blood meals tend to defecate quicker than those
that have ingested smaller amounts [43]. Thus, the vector-
ial capacity of T. infestans might diminish less than that of
P. megistus and T. brasiliensis as the population density
increases because there is less of a reduction in bloodmeal
size for T. infestans.
Concluding remarks
We propose that the great capacity for exploiting blood
resources from available hosts in dwellings enabled T.
infestans to spread and become the most important vector
of T. cruzi in the south cone of South America. This
proposal is in accordance with the central role of the
insect–host interactions in the biology and vectorial capa-
city of Triatominae.
Brazil is winning the battle against T. infestans. How-
ever, other indigenous species (especially T. brasiliensis
and P. megistus) are capable of invading and colonizing
houses. These ubiquitous species have been found in nat-
ural and artificial sites close to human dwellings and are
associated with wild or peridomestic animals. In this
scenario, it is important to understand how the features
of the hosts (physiology, blood nutritional value and groom-
ing behavior) frequently used as food sources interfere with
Triatominae population dynamics to modulate its spatial
distribution and predisposition to spread.
Acknowledgements
We thank C.J. Schofield for reviewing the manuscript. We also thank
Bruce Alexander, Ricardo N. Araujo and Adriana Santos for their
comments. This work was supported by CNPq, FAPEMIG, CPqRR/
FIOCRUZ and benefited from international collaboration through the
ECLAT network.
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